Give What You Get: Capuchin Monkeys (Cebus apella)

and 4-Year-Old Children Pay Forward Positive and

Negative Outcomes to Conspecifics
Kristin L. Leimgruber1"*, Adrian F. Ward2", Jane Widness1, Michael I. Norton3, Kristina R. Olson1,
Kurt Gray4, Laurie R. Santos1
1 Department of Psychology, Yale University, New Haven, Connecticut, United States of America, 2 Department of Psychology, Harvard University, Cambridge,
Massachusetts, United States of America, 3 Harvard Business School, Harvard University, Boston, Massachusetts, United States of America, 4 Department of Psychology,
University of Maryland, College Park, Maryland, United States of America

Abstract
The breadth of human generosity is unparalleled in the natural world, and much research has explored the mechanisms
underlying and motivating human prosocial behavior. Recent work has focused on the spread of prosocial behavior within
groups through paying-it-forward, a case of human prosociality in which a recipient of generosity pays a good deed forward
to a third individual, rather than back to the original source of generosity. While research shows that human adults do
indeed pay forward generosity, little is known about the origins of this behavior. Here, we show that both capuchin
monkeys (Cebus apella) and 4-year-old children pay forward positive and negative outcomes in an identical testing
paradigm. These results suggest that a cognitively simple mechanism present early in phylogeny and ontogeny leads to
paying forward positive, as well as negative, outcomes.

Citation: Leimgruber KL, Ward AF, Widness J, Norton MI, Olson KR, et al. (2014) Give What You Get: Capuchin Monkeys (Cebus apella) and 4-Year-Old Children Pay
Forward Positive and Negative Outcomes to Conspecifics. PLoS ONE 9(1): e87035. doi:10.1371/journal.pone.0087035
Editor: Roscoe Stanyon, University of Florence, Italy
Received July 1, 2013; Accepted December 17, 2013; Published January 29, 2014
Copyright: ß 2014 Leimgruber et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Funding: This research was supported in part by funding from The Templeton Positive Neuroscience Award and the National Science Foundation Research
Experiences for Undergraduates Program (SMA-1004797). The funders had no role in study design, data collection and analysis, decision to publish, or preparation
of the manuscript.
Competing Interests: Laurie R. Santos is a PLOS ONE Editorial Board member but this does not cause a conflict of interest, nor does this alter the authors’
adherence to all the PLOS ONE policies on sharing data and materials.
* E-mail: Kleimgruber@gmail.com
"
KL and AW are joint first authors

Introduction
Humans frequently and willingly engage in costly behaviors that
benefit others, even when their actions are anonymous and when
those helped are total strangers [1,2]. This proclivity for prosocial
behavior is unparalleled in the natural world and is thus thought to
play a key role in large-scale cooperation unique to human society
[3]. As a result, much research has been devoted to understanding
the social, cognitive, and biological processes that encourage (and
discourage) prosociality in adult humans [4–7]. Recently,
researchers have begun to explore the spread of prosocial behavior
within populations [8,9]; specifically, several studies have exam-
ined when and why people pay forward prosocial behavior
[10,11]. This concept of ‘‘paying-it-forward’’ is simple: Person A
helps Person B and Person B, rather than paying this kindness back
to Person A, pays it forward to Person C, thus facilitating the spread
of prosocial behavior beyond the dyad to a larger group of
individuals. While experimental research [9,12,13] and real-life
accounts [14] indicate that humans do pay forward positive
outcomes, the psychological underpinnings of such behavior
remain unresolved. Traditional explanations for paying forward
positive outcomes tend to rely on socially and cognitively complex
mechanisms including gratitude [15–18], cultural and moral
norms [19,20], and processes requiring sophisticated perspective-
taking abilities [21]. Taken together, these social and cognitive
constraints might suggest that paying forward generosity is a
uniquely human phenomenon.
However, a comprehensive review of the existing literature
suggests that the tendency to pay-it-forward may instead be
explained by more rudimentary behavioral strategies that are not,
in fact, unique to human adults. Specifically, it is possible that
people act on the basis of the maxim: ‘‘help anyone, if helped by
someone’’ (hereafter, help-if-helped) [22]. Unlike more cognitively
complex explanations for the propagation of prosocial behavior,
this strategy does not require memory of the identities of
interaction partners [23], sensitivity to one’s own reputational
status [24,25], the capacity to calculate the potential costs and
benefits of prosocial behavior [26], or the use of self-control to
inhibit initial selfish urges [27,28]; instead, the strategy simply
requires that individuals do to others what was done to them. Both
mathematical models [29,30] and laboratory simulations
[12,13,31] have demonstrated that a simple rule like help-if-
helped could lead to self-sustaining pay-it-forward systems.
Moreover, experiments indicating that rats (Rattus norvegicus) pay
forward helping behaviors [32] provide further evidence that
complex and/or uniquely human social and cognitive capacities
are not required for organisms to pay forward generosity. Indeed,
these findings show that a help-if-helped strategy is not only

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sufficient to support the propagation of prosocial behavior within
populations, but also that it likely predates more discriminating
forms of cooperative behavior that rely upon the complex social
and cognitive abilities found only in human adults.
Furthermore, the majority of existing studies investigating the
psychology of paying-it-forward focus exclusively on the prosocial
side of paying behavior forward–that is, on paying forward positive
outcomes. However, laboratory simulations of pay-it-forward
behavior suggest that negative outcomes are just as likely to be
paid forward in public goods games as positive ones [9], and
experimental evidence suggests that–in some situations–adults pay
forward greed more than generosity [10]. These findings, along
with a long history of literature on displaced aggression [33], call
into question the proposed role of prosocial intentions, positive
emotions and moral norms in paying forward like outcomes.
Instead, they suggest the existence of a strategy even more simple
than help-if-helped: they suggest that pay-it-forward behavior may
be based on the rudimentary rule of ‘‘give what you get’’
(hereafter, give-what-you-get).
Taken as a whole, this set of findings hints that existing
research–which typically divides paying-it-forward into separate
positive and negative phenomena–may be neglecting a more
parsimonious explanation for the propagation of behavior in
general. While some accounts of paying it forward favor
cognitively and morally rich accounts of human kindness
[34,35], empirical evidence suggests that these behaviors may
instead be rooted in a general tendency to reciprocate both
positive and negative behaviors in kind [9]. If this simple
explanation holds true, we would expect to see behaviors
consistent with a give-what-you-get mechanism present early in
human development, and possibly even in non-human primates.
The current study tests this possibility by examining pay-it-
forward tendencies in 4-year-old children and capuchin monkeys
(Cebus apella). While there is evidence that capuchin monkeys [36–
38] and young children [39–42] consistently take advantage of no-
cost opportunities to act prosocially toward conspecifics, both
groups lack certain capacities key to current explanations of
paying-it-forward in human adults. Specifically, capuchin monkeys
largely fail at tasks that rely on perspective-taking abilities [43–47],
self-awareness [48,49], and the ability to evaluate and reflect upon
their own knowledge states [50,51] – all cognitive capacities
assumed necessary for the experience of gratitude [52–54] and
implicated in current explanations for paying forward generosity.
Similarly, before the age of five, children have difficulty in
evaluating the perspectives and knowledge states of others in a
consistent manner [55–58] and in evaluating and reflecting upon
their own thoughts and knowledge states [59]. Unlike capuchin
monkeys, however, young children have likely been exposed to
social and moral norms advocating paying forward generosity in
some form or another. Testing these populations using an identical
paradigm allows us to identify the minimal cognitive abilities
required to pay-it-forward and illuminates the role uniquely
human social and moral norms play in the propagation of paying
forward generosity.
Participants in the current study took part in a chain of non-
anonymous donation games in which individuals first received a
positive or negative outcome from a member of their social group,
and then had the chance to distribute a positive or negative
outcome to a different member of this social group. We used only
‘‘no-cost’’ options, in which participants making donation
decisions received the same outcome regardless of the outcome
they chose to deliver to a group member. The use of a ‘‘no-cost,’’
(or non-zero-sum) paradigm reduces the role of self-interested
motivations, accounts for between-species differences in self-
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Give What You Get
control and/or reputational concerns, and minimizes cognitive
demands imposed by trade-off related calculations. Controlling for
these factors allowed us to explore the minimal social and
cognitive factors underlying pay-it-forward strategies, thus making
it possible to identify the most parsimonious explanation for the
donation behaviors observed in monkeys and children.
Materials and Methods
Ethics Statement
This study was carried out in strict accordance with the
recommendations in the Guide for Care and Use of Laboratory
Animals of the National Institutes of Health. The protocol for non-
human primates was approved by the Institutional Animal Care
and Use Committee at Yale University (Protocol Number: #2008-
10678). The treatment of human participants in studies described
in this paper was in accordance with the ethical standards of the
American Psychological Association. Participants’ parents provid-
ed written informed consent and all procedures were approved by
the Human Research Protection Program at Yale University.
Participants
Monkey participants were 4 brown capuchins (Cebus apella)
ranging in age from 5–15 years at the conclusion of the study (1
male [NN], 3 females [HG, HR, JM]; Mage = 134.86 months;
SD = 55.05). Our capuchin participants were members of the
Yale Comparative Cognition Laboratory colony where they
were socially housed in a large indoor enclosure equipped with
natural branches and toys. Capuchins were fed monkey chow
prior to testing and had access to water ad libitum. All
participants had previous experience with reward distribution
tasks involving conspecifics [36] and were familiar with one
another prior to testing. To control for the effects of previous
experience and developmental differences in social cognitive
abilities, only mature adult monkeys who had previously
demonstrated an understanding of the apparatus (Unpublished
data) were involved in the current study. Although these strict
selection criteria limited the number of monkeys we were able
to include in the study, our final sample size is nonetheless
comparable to those in other studies of social cognition in
brown capuchin monkeys [38,60–62].
We also tested 31 four-year-old children (10 males, 21 females;
Mage = 54.68 months; SD = 3.45) recruited from preschools in the
greater New England area. Participants were tested in mixed
gender groups comprised of children from the same classroom; as
a result, all children were familiar with one another prior to
testing. Care was taken to ensure that children never received
from, or gave to, members of their own family. Group size was
constrained by the number of consenting participants per class,
with groups ranging in size from 3–7 individuals.
General Methods
Testing was performed using identical novel apparatuses for
the monkeys (Figure 1) and the children (Figure 2) that allowed
participants to choose between two distinct distributions. Each
distribution provided an allocation for an Actor (the participant
manipulating the apparatus), and an allocation for a Recipient
(a second participant who merely received whatever he/she was
given). The apparatus was situated between the Actor and the
Recipient such that the two participants were able to see one
another and the distribution options over the top of the
apparatus. In order to equate the non-verbal methods as closely
as possible across the two populations, children were asked not
to speak to one another or signal their preferences in any way.
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The Actor was always the participant seated on the side of the
apparatus with two identical levers. By pulling the lever on her
left, the Actor could distribute the leftmost allocations to herself
and the Recipient; by pulling the lever on her right, the Actor
could distribute the rightmost allocations to herself and the
Recipient. Allocations were simultaneously delivered via a chute
to both participants immediately following the Actor’s choice,
and the two remaining allocations were removed from the
apparatus by the experimenter.
Using this apparatus, monkeys and children participated in a
series of overlapping donation games, such that each participant in
a chain first received from, and then gave to different conspecific
members of their social groups. Test sessions began when an initial
Actor distributed an outcome to a conspecific Recipient. After
both participants had collected their respective allocations, the
Recipient moved to the opposite side of the apparatus to assume
the role of Actor and the previous Actor left the testing area. At
this point, a third conspecific–ignorant to the outcome of the
previous interaction–entered the testing area to assume the role of
Recipient, and the new Actor was asked to choose between the
same distribution options presented to the previous Actor. This
process continued until all participants had received from, and
subsequently given to, a conspecific group member, with the initial
Actor serving as the recipient for the final participant. Data from
Figure 1. Testing apparatus used for monkeys. Monkey Actors pulled one of the two levers to choose an outcome to distribute to the Receiver
situated on the other side of the apparatus.
doi:10.1371/journal.pone.0087035.g001
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Give What You Get
the initial Actor was excluded from analysis. Efforts were made to
ensure that testing and data collection procedures were identical
between species whenever possible (however, see Methods S1,
Table S1, and Table S2 for between-species methodological
differences).
Allocations were placed inside of clear, round, plastic
containers that allowed for easy distribution via the apparatus.
The placement of the distribution options (positive/negative)
into the apparatus was counterbalanced to control for the
possible role of side biases in participant’s donation choices (see
Methods S1, Table S1, and Table S2 for more details). For
each test trial, Actors had the option to deliver one of two
outcomes to the Recipient: a positive outcome that delivered a
high-value allocation to both herself and the recipient, or a
negative outcome that delivered a high-value allocation to
herself and a low-value allocation to the Recipient. A positive
outcome for monkeys consisted of a grape for both the Actor
and the Recipient; a negative outcome consisted of a grape for
the Actor and a piece of spinach for the Recipient. A positive
outcome for children consisted of 4 small, star-shaped stickers
for both the Actor and Recipient; a negative outcome consisted
of 4 small, star-shaped stickers for the Actor and 1 small, star-
shaped sticker for the Recipient. Actors always received the
high-value reward, regardless of what they chose to distribute to
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Figure 2. Testing apparatus used for children. Actors pulled one of the two levers to choose an outcome to distribute to the Receiver situated
on the other side of the apparatus.
doi:10.1371/journal.pone.0087035.g002
Receivers; thus, there was no cost to generosity and no benefit
from greed–participants’ distributions to conspecifics revealed
the tendency to pay forward outcomes, divorced from potential
selfish motives present in zero-sum distribution tasks. In
addition, using the same value reward across both options for
the Actors removed any confounds related to differences in or
distractions from their own outcome.
Results
Actors’ distributions were strongly related to previously received
outcomes, for both monkeys (n = 4 participants, 22 trials, Fisher’s
exact, p = .03) and children (n = 48 children, 48 trials, Fisher’s
exact, p = .009). Monkeys paid forward negative outcomes 75% of
the time and positive outcomes 80% of the time; children paid
forward negative outcomes 72% of the time and positive outcomes
70% of the time. The rates at which positive versus negative
outcomes were paid forward did not significantly differ in monkeys
(X2(1, N = 17) = .06, p = .81) or children (X2(1, N = 34) = .12,
p = .73). Similarly, the rates at which children versus monkeys
paid forward positive (X2(1, N = 33) = .38, p = .54) and negative
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Give What You Get
(X2(1, N = 37) = .04, p = .85) outcomes were not statistically
different across species. See Figure 3 for all results.
We also performed a series of logistic regressions in order to
confirm that the giving behavior of both children and monkeys
truly reflected a tendency to pay forward both positive and
negative behavior in kind. If this was the case, then previously
received outcomes should predict giving behavior even when
controlling for all other possible variables (e.g., identity of initial
Actor, identity of Recipient). A regression on monkey giving
behavior (positive, negative) using the predictors of initial Actor
identity, focal participant identity, final Receiver identity, and received outcome
(positive, negative) revealed that only received outcome affected giving
behavior, Wald’s X2(1) = 7.34, p,.01; all other predictors p..45. A
regression on children’s giving behavior (positive, negative) using
‘the predictors of initial Actor gender, focal participant gender, final
Receiver gender, and received outcome (positive, negative) revealed that
only received outcome affected giving behavior, Wald’s X2(1) = 7.34,
p,.01; all other predictors p..16. Taken together, these analyses
confirm that the giving behavior of both children and monkeys can
be attributed to previously received outcomes–that is, they paid
both positive and negative behavior forward in kind. See Tables
S1 and S2 for trial-by-trial data.
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Figure 3. Percentage of total trials in which monkeys and children paid forward positive and negative outcomes after receiving
positive and negative outcomes.
doi:10.1371/journal.pone.0087035.g003
Discussion
Our results suggest that the propagation of both positive and
negative behavior within social groups may stem from a
mechanism that is both cognitively simple and evolutionarily
old. Our finding that monkeys and 4-year-old children paid
forward positive outcomes to conspecifics is line with previous
behavioral findings in rats [32], and confirms that the act of paying
forward positive events does not require complex emotions [15],
human-specific norms [20], or sophisticated perspective-taking
abilities [21]. Our results also indicate that both populations paid
forward positive and negative outcomes, demonstrating that paying
forward behavior is not limited to prosocial interactions. Instead,
our findings suggest that paying forward behavior in monkeys and
children is best explained by a simple give-what-you-get mecha-
nism–one that may be the foundation upon which more complex
cooperative behaviors are built in adult humans.
Overall, our findings are consistent with a contingency-based
give-what-you-get strategy, a form of generalized reciprocity in
which like begets like, regardless of the specific recipient or the
valence of the outcome [23]. Giving what you get is less cognitively
complex than other forms of reciprocity, and so is a likely
explanation for group-level cooperation in non-human animals
[63,64]. Importantly, because contingency-based strategies like
give-what-you-get are not sensitive to recipient identity, individ-
uals employing them need not differentiate between paying
outcomes forward and paying outcomes back; they are simply
motivated to reciprocate outcomes in kind. As a result, this
explanation implies that the pattern of behavior of monkeys and
children in the current study may not necessarily be specific to
paying-it-forward, but rather a reflection of a motivation to
reciprocate outcomes in general. If this were the case, we would
expect similar patterns of giving whether the recipient was the
same or different than the individual from whom an allocation was
received; that is, we would expect minimal differences between
paying behavior forward and paying behavior back. However, if
the mechanism underlying our results is specific to paying-it-
forward, we would expect different patterns of giving when
individuals have the opportunity to pay outcomes back to the initial
actor. While the current data do not allow us to make this
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Give What You Get
distinction, future research could examine the extent to which
common and distinct mechanisms underlie these two related
behaviors.
In our results, monkeys and children appear to pay forward like
outcomes at equally high rates, despite evidence suggesting
predispositions toward prosociality in other contexts [39–43].
The current data do not include baseline levels of giving, but a
follow-up study comparing general rates of positive/negative
outcome distribution with rates after participants receive a
positive/negative outcome themselves would offer insight into
the relative strength of the drive to pay forward positive vs.
negative outcomes. Nonetheless, our findings show a clear pattern
of behavior in which giving in both monkeys and children is
influenced by the valence of received outcomes; these data suggest
that a ‘‘give what you get’’ strategy drives the tendency to pay
forward both positive and negative outcomes.
A second (and not mutually exclusive) possibility consistent with
our results is that pay-it-forward tendencies are driven by basic
affective processes, ones that may be precursors to the more
sophisticated emotions observed in adult humans [10]. Whereas
affect (i.e. positive and negative feelings) occur automatically
[65,66] and across species [67], gratitude is considered a
secondary emotion requiring additional cognitive resources to
interpret initial basic affective responses [15,68,69]. Capuchin
monkeys do not possess the suite of cognitive abilities associated
with explanations relying on complex emotions like gratitude, and
thus these emotions cannot drive pay forward behaviors as some
have hypothesized [15–18]. However, both children and capu-
chins possess basic affective processes that may drive paying
forward of both positive and negative outcomes. Indeed, affect has
been shown to motivate future behavior in adult humans
[10,31,70–73] as well other primate species [74–76]; further
research may determine the role of affective factors in pay-it-
forward behaviors in capuchin monkeys and children.
While the form of paying-it-forward we observed in capuchin
monkeys and young children does not require secondary emotions,
perspective-taking abilities, or uniquely human social or moral
norms, these factors are likely important in adult humans. Our
results therefore hint that the mechanism underlying pay-it-
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forward behavior in children and monkeys serves as a framework
around which more sophisticated social, emotional, and moral
decision-making systems are built in adult humans. For example,
while adult humans–like monkeys and children–surely experience
basic affective responses after experiencing a positive or negative
outcome, they may be uniquely be able to draw upon a more
sophisticated suite of cognitive abilities with which to reflect upon
their experiences and determine their future actions. Whereas
monkeys and children in our study consistently paid forward
negative outcomes despite the no-cost nature of the task, the
general tendency to pay forward negative outcomes in adults may
be overshadowed by more cognitively sophisticated processes such
as cost/benefit analyses [26] or concerns about maintaining one’s
positive reputation within the group [77–79]. Indeed, research
shows that human adults often act in strikingly self-interested ways
when cost/benefit analyses tip in their favor and the likelihood of
negative reputational repercussions is low [80–82], suggesting that
increased cognitive sophistication doesn’t necessitate increased
prosociality, but rather facilitates flexible decision-making pro-
cesses. Likewise, it is probable that paying forward generosity in
human adults is not merely due to general positive affect, but is
instead the result of further cost/benefit analyses, secondary
emotions like gratitude [15], adherence to cultural or religious
norms [19], and/or means to attain the warm glow that comes as
the result of being the cause of another’s good fortune [80]. In the
end, though, all of these more cognitively complex factors may
merely be building on–or modifying–the simple strategies evident
in the behavior of organisms such as capuchin monkeys and
human children.
Our results indicate that the propagation of prosocial behavior
within groups is not rooted in prosocial motives alone, but instead
emerges via a simple mechanism, shared across phylogeny and
ontogeny, that encourages paying forward both positive and
negative behaviors in kind. Our results suggest that even the most
heartwarming acts of paying forward generosity likely have their
roots in a simple mechanism that is not limited to prosocial
tendencies. While emotions like gratitude and uniquely human
norms likely play a role in the extraordinary cases of paying
forward generosity that make newspaper headlines, our data
References
1. Penner LA, Dovidio JF, Piliavin JA, Schroeder DA (2005) Prosocial Behavior:
Multilevel Perspectives. Annu Rev Psychol 56: 365–392.
2. Henrich J, Bowles S, Boyd RT, Opfensitz A, Richerson PJ, et al. (2003) The
cultural and genetic evolution of human cooperation. In: Hammerstein P, editor.
Genetic and Cultural Evolution of Cooperation. MIT Press. 445–468. Available:
http://books.google.com/books?hl = en&lr = &id = aVh9jtWbG0wC&pgis = 1.
Accessed 12 October 2013.
3. Jaeggi AV, Burkart JM, Van Schaik CP (2010) On the psychology of
cooperation in humans and other primates: Combining the natural history
and experimental evidence of prosociality. Philos Trans R Soc Lond B Biol Sci
365: 2723–2735. Available: http://www.pubmedcentral.nih.gov/articlerender.
fcgi?artid = 2936168&tool = pmcentrez&rendertype = abstract. Accessed 2012
March 14.
4. Batson CD, Powell AA (2003) Altruism and prosocial behavior. In: Millon T,
Lerner MJ, editors. Handbook of psychology: Personality and Social Psychology.
Hoboken, NJ: John Wiley & Sons, Inc. 463–484.
5. Rilling JK, Sanfey AG (2011) The neuroscience of social decision-making. Annu
Rev Psychol 62: 23–48. Available: http://www.annualreviews.org/doi/abs/10.
1146/annurev.psych.121208.131647. Accessed 2013 September 20.
6. Bekkers R, Wiepking P (2010) A Literature Review of Empirical Studies of
Philanthropy: Eight Mechanisms That Drive Charitable Giving. Nonprofit
Volunt Sect Q 40: 924–973. Available: http://nvs.sagepub.com/content/40/5/
924.short. Accessed 2013 October 12.
7. MacDonald K, MacDonald TM (2010) The Peptide That Binds: A Systematic
Review of Oxytocin and its Prosocial Effects in Humans. Available: http://
informahealthcare.com/doi/abs/10.3109/10673220903523615. Accessed 2013
October 12.
8. Rand DG, Arbesman S, Christakis NA (2011) Dynamic social networks promote
cooperation in experiments with humans. Proc Natl Acad Sci U S A 108:
PLOS ONE | www.plosone.org
Give What You Get
suggest paying-it-forward may propagate and persist within social
groups, even in the absence of these factors. Although a fascination
with the propagation of kindness–and a tendency to explain these
behaviors in moralistic terms–may be uniquely human, the
mechanism underlying this behavior is likely not.
Supporting Information
Table S1 Positive/Negative outcome distribution patterns
within testing chains for monkeys. Each row represents a discrete
test session; monkeys only participated in one test session per day.
Trials in which monkeys ‘gave what they got’ are bolded. Trials in
which monkeys paid forward negative outcomes are highlighted in
blue; trials in which monkeys paid forward positive outcomes are
highlighted in yellow.
(TIF)
Table S2 Positive/Negative outcome distribution patterns
within testing chains for children. Each row represents a single
testing chain; variance in chain length is due to variance in the
number of consenting children per classroom. Trials in which
children (males = M; females = F) ‘gave what they got’ are bolded.
Trials in which children paid forward negative outcomes are
highlighted in blue; trials in which children paid forward positive
outcomes are highlighted in yellow.
(TIF)
Methods S1 Supplemental methods.
(DOCX)
Acknowledgments
The authors wish to thank Anna Merrill, Nick Buttrick, Melissa Baranay,
Andre Alexander, Sarah Arn, Lara Boyle, Channler Hill, Adam Hoffman,
and Kelly Reina for their help in running these studies.
Author Contributions
Conceived and designed the experiments: KLL AFW MIN KRO KG
LRS. Performed the experiments: KLL JW. Analyzed the data: KLL
AFW. Wrote the paper: KLL AFW MIN KRO KG LRS.
19193–19198. Available: http://www.pnas.org/content/108/48/19193. Ac-
cessed 2013 October 18.
9. Fowler JH, Christakis NA (2010) Cooperative behavior cascades in human social
networks. Proc Natl Acad Sci U S A 107: 5334–5338. Available: http://www.
pnas.org/content/107/12/5334.short. Accessed 2013 September 20.
10. Gray K, Ward AF, Norton MI (2012) Paying It Forward: Generalized
Reciprocity and the Limits of Generosity. J Exp Psychol Gen 142: 1–8.
Available: http://www.ncbi.nlm.nih.gov/pubmed/23244034. Accessed 2013
March 25.
11. Grant A, Dutton J (2012) Beneficiary or benefactor: are people more prosocial
when they reflect on receiving or giving? Psychol Sci 23: 1033–1039. Available:
http://pss.sagepub.com/content/23/9/1033.abstract. Accessed 2013 October
18.
12. Greiner B, Levati MV (2005) Indirect reciprocity in cyclical networks: An
experimental study. J Econ Psychol 26: 711–731. Available: http://linkinghub.
elsevier.com/retrieve/pii/S0167487005000334. Accessed 2011 September 2.
13. Stanca L (2009) Measuring indirect reciprocity: Whose back do we scratch? J
Econ Psychol 30: 190–202. Available: http://linkinghub.elsevier.com/retrieve/
pii/S016748700800072X. Accessed 2011 August 8.
14. Lotze K (2012) Pay It Forward Wedding Dress. Huffingt Post. Available: http://
www.huffingtonpost.com/tag/pay-it-forward-wedding-dress. Accessed 2013 Oc-
tober 13.
15. Bartlett MY, DeSteno D (2006) Gratitude and prosocial behavior: Helping when
it costs you. Psychol Sci 17: 319–325. Available: http://www.ncbi.nlm.nih.gov/
pubmed/16623689.
"
KL and AW are joint first authors
6 January 2014 | Volume 9 | Issue 1 | e87035

16. Nowak MA, Roch S (2007) Upstream reciprocity and the evolution of gratitude.
Proc Biol Sci 274: 605–609. Available: http://www.pubmedcentral.nih.gov/
articlerender.fcgi?artid = 2197219&tool = pmcentrez&rendertype = abstract. Ac-
cessed 2011 July 26.
17. Tsang JA (2006) BRIEF REPORT Gratitude and prosocial behaviour: An
experimental test of gratitude. Cogn Emot 20: 138–148. Available: http://www.
tandfonline.com/doi/abs/10.1080/02699930500172341. Accessed 2013 March
18.
18. Tsang JA (2007) Gratitude for small and large favors: A behavioral test. J Posit
Psychol 2: 157–167. Available: http://www.tandfonline.com/doi/abs/10.1080/
17439760701229019. Accessed 2013 May 2.
19. Emmons RA, Crumpler CA (2000) Gratitude as a Human Strength: Appraising
the Evidence. J Soc Clin Psychol 19: 56–69. Available: http://guilfordjournals.
com/doi/abs/10.1521/jscp.2000.19.1.56. Accessed 2013 October 11.
20. Nowak MA, Sigmund K (2005) Evolution of indirect reciprocity. Nature 437:
1291–1298. Available: http://www.ncbi.nlm.nih.gov/pubmed/16251955. Ac-
cessed 2011 July 15.
21. Whiten A, Bryne R (1997) Machiavellian Intelligence II: Extensions and
Evaluations Cambridge University Press. p. 403. Available: http://books.google.
com/books?hl = en&lr = &id = bV9yeFV6_ckC&pgis = 1. Accessed 2013 Octo-
ber 11.
22. Van Doorn GS, Taborsky M (2012) The evolution of generalized reciprocity on
social interaction networks. Evolution 66: 651–664. Available: http://www.ncbi.
nlm.nih.gov/pubmed/22380430. Accessed 2013 October 11.
23. Trivers R (1971) The Evolution of Reciprocal Altruism. Q Rev Biol. Available:
http://www.jstor.org/stable/10.2307/2822435. Accessed 2013 October 11.
24. Mohtashemi M, Mui L (2003) Evolution of indirect reciprocity by social
information: The role of trust and reputation in evolution of altruism. J Theor
Biol 223: 523–531. Available: http://www.sciencedirect.com/science/article/
pii/S0022519303001437. Accessed 2013 October 11.
25. Ohtsuki H, Iwasa Y (2004) How should we define goodness?–Reputation
dynamics in indirect reciprocity. J Theor Biol 231: 107–120. Available: http://
www.sciencedirect.com/science/article/pii/S0022519304002772. Accessed
2013 October 11.
26. Lehmann L, Keller L (2006) The evolution of cooperation and altruism–a
general framework and a classification of models. J Evol Biol 19: 1365–1376.
Available: http://www.ncbi.nlm.nih.gov/pubmed/16910958. Accessed 2013
September 17.
27. Fehr E, Schmidt KM (1999) A Theory Of Fairness, Competition, and
Cooperation. Q J Econ 114: 817–868. Available: http://www.
mitpressjournals.org/doi/abs/10.1162/003355399556151.
28. Nowak MA, Page KM, Sigmund K (2000) Fairness versus reason in the
ultimatum game. Science (80- ) 289: 1773–1775. Available: http://www.ncbi.
nlm.nih.gov/pubmed/10976075. Accessed 2013 September 26.
29. Barta Z, McNamara JM, Huszár DB, Taborsky M (2011) Cooperation among
non-relatives evolves by state-dependent generalized reciprocity. Proc Biol Sci
278: 843–848. Available: http://rspb.royalsocietypublishing.org/content/278/
1707/843.short. Accessed 2012 April 18.
30. Iwagami A, Masuda N (2010) Upstream reciprocity in heterogeneous networks. J
Theor Biol 265: 297–305. Available: http://www.sciencedirect.com/science/
article/pii/S0022519310002390. Accessed 2011 November 21.
31. Dufwenberg M, Gneezy U, Güth W, Van Damme E (2000) An experimental
test of direct and indirect reciprocity in case of complete and incomplete
information. Available: http://www.econstor.eu/handle/10419/62163. Ac-
cessed 2013 October 11.
32. Rutte C, Taborsky M (2007) The influence of social experience on cooperative
behaviour of rats (Rattus norvegicus): Direct vs generalised reciprocity. Behav
Ecol Sociobiol 62: 499–505. Available: http://www.springerlink.com/index/10.
1007/s00265-007-0474-3. Accessed 2012 March 11.
33. Marcus-Newhall A, Pedersen WC, Carlson M, Miller N (2000) Displaced
aggression is alive and well: A meta-analytic review. J Pers Soc Psychol 78: 670–
689. Available: http://www.ncbi.nlm.nih.gov/pubmed/10794373. Accessed
2013 October 11.
34. Emerson RW (1841) Compensation. Essays: First and Second Series. Gloucester,
UK: Dodo Press.
35. Franklin B (1784) Letter to Benjamin Webb. The Works of Benjamin Franklin.
Boston: Hilliard, Gray, and Company.
36. Lakshminarayanan VR, Santos LR (2008) Capuchin monkeys are sensitive to
others’ welfare. Curr Biol 18: R999–R1000. Available: http://www.
sciencedirect.com/science/article/pii/S0960982208011354. Accessed 2013 Oc-
tober 11.
37. de Waal FBM, Leimgruber K, Greenberg AR (2008) Giving is self-rewarding for
monkeys. Proc Natl Acad Sci U S A 105: 13685–13689. Available: http://www.
pnas.org/content/105/36/13685.short. Accessed 2013 September 22.
38. Takimoto A, Kuroshima H, Fujita K (2010) Capuchin monkeys (Cebus apella)
are sensitive to others’ reward: An experimental analysis of food-choice for
conspecifics. Anim Cogn 13: 249–261. Available: http://www.ncbi.nlm.nih.
gov/pubmed/19609580. Accessed 2011 October 27.
39. Blake PR, Rand DG (2010) Currency value moderates equity preference among
young children. Evol Hum Behav 31: 210–218. Available: http://linkinghub.
elsevier.com/retrieve/pii/S1090513809000658. Accessed 2011 June 9.
40. Buhrmester D, Goldfarb J, Cantrell D (1992) Self-Presentation when Sharing
with Friends and Nonfriends. J Early Adolesc 12: 61–79. Available: http://jea.
sagepub.com/cgi/doi/10.1177/0272431692012001004. Accessed 2011 June 9.
PLOS ONE | www.plosone.org
Give What You Get
41. Gummerum M, Hanoch Y, Keller M, Parsons K, Hummel A (2010)
Preschoolers’ allocations in the dictator game: The role of moral emotions. J
Econ Psychol 31: 25–34. Available: http://linkinghub.elsevier.com/retrieve/
pii/S0167487009000993. Accessed 2011 September 11.
42. Harbaugh W, Krause K (2000) Children’s altruism in public good and dictator
experiments. Econ Inq 38: 95–109. Available: http://doi.wiley.com/10.1111/j.
1465-7295.2000.tb00006.x. Accessed 2013 October 11.
43. Anderson JR, Sallaberry P, Barbier H (1995) Use of experimenter-given cues
during object-choice tasks by capuchin monkeys. Anim Behav 49: 201–208.
Available: http://www.sciencedirect.com/science/article/pii/
0003347295801685. Accessed 2013 October 11.
44. Barnes JL, Hill T, Langer M, Martinez M, Santos LR (2008) Helping behaviour
and regard for others in capuchin monkeys (Cebus apella). Biol Lett 4: 638–640.
Available: http://rsbl.royalsocietypublishing.org/content/4/6/638.short. Ac-
cessed 2012 March 30.
45. Hare B, Addessi E, Call J, Tomasello M, Visalberghi E (2003) Do capuchin
monkeys, Cebus apella, know what conspecifics do and do not see? Anim Behav
65: 131–142. Available: http://www.sciencedirect.com/science/article/pii/
S0003347202920178. Accessed 2013 October 11.
46. Itakura S, Anderson JR (1996) Learning to use experimenter-given cues during
an object-choice task by a capuchin monkey. Curr Psychol Cogn 15: 103–112.
47. Hattori Y, Kuroshima H, Fujita K (2010) Tufted capuchin monkeys (Cebus
apella) show understanding of human attentional states when requesting food
held by a human. Anim Cogn 13: 87–92. Available: http://www.ncbi.nlm.nih.
gov/pubmed/19517147. Accessed 2013 October 11.
48. Anderson JR, Roeder JJ (1989) Responses of capuchin monkeys (Cebus apella)
to different conditions of mirror-image stimulation. Primates 30: 581–587.
Available: http://link.springer.com/10.1007/BF02380884. Accessed 2013 Oc-
tober 11.
49. de Waal FBM, Dindo M, Freeman CA, Hall MJ (2005) The monkey in the
mirror: Hardly a stranger. Proc Natl Acad Sci U S A 102: 11140–11147.
Available: http://www.pnas.org/content/102/32/11140.short. Accessed 2013
September 22.
50. Beran MJ, Smith JD, Coutinho MVC, Couchman JJ, Boomer J (2009) The
psychological organization of ‘‘uncertainty’’ responses and ‘‘middle’’ responses:
A dissociation in capuchin monkeys (Cebus apella). J Exp Psychol Anim Behav
Process 35: 371–381. Available: http://www.ncbi.nlm.nih.gov/pubmed/
19594282. Accessed 2013 October 14.
51. Paukner A, Anderson JR, Fujita K (2006) Redundant food searches by capuchin
monkeys (Cebus apella): A failure of metacognition? Anim Cogn 9: 110–117.
Available: http://www.ncbi.nlm.nih.gov/pubmed/16184375. Accessed 2013
October 11.
52. Ben-Ze’Ev A, Oatley K (1996) The Intentional and Social Nature of Human
Emotions: Reconsideration of the Distinction Between Basic and Non-basic
Emotions. J Theory Soc Behav 26: 81–94. Available: http://doi.wiley.com/10.
1111/j.1468-5914.1996.tb00287.x.
53. Gallup GG (1998) Self-awareness and the evolution of social intelligence. Behav
Processes 42: 239–247. Available: http://www.sciencedirect.com/science/
article/pii/S037663579700079X. Accessed 2013 October 11.
54. McAdams DP, Bauer JJ (2004) Gratitude in Modern Life: Its Manifestations and
Development. In: Emmons RA, Mccullough ME, editors. The Psychology of
Gratitude: Series in Affective Science. New York, NY: Oxford University Press.
81–99.
55. Epley N, Caruso E, Bazerman MH (2006) When perspective taking increases
taking: Reactive egoism in social interaction. J Pers Soc Psychol 91: 872–889.
Available: http://www.ncbi.nlm.nih.gov/pubmed/17059307.
56. Perner J (1991) Understanding the representational mind. Learning, develop-
ment, and conceptual change. Cambridge, MA: The MIT Press.
57. Wellman HM, Cross D, Watson J (2001) Meta-analysis of theory-of-mind
development: The truth about false belief. Child Dev 72: 655–684. Available:
http://www.ncbi.nlm.nih.gov/pubmed/11405571. Accessed 2013 September
23.
58. Wimmer H, Perner J (1983) Beliefs about beliefs: Representation and
constraining function of wrong beliefs in young children’s understanding of
deception. Cognition 13: 103–128. Available: http://www.sciencedirect.com/
science/article/pii/0010027783900045. Accessed 2013 October 11.
59. Flavell JH, Green FL, Flavell ER (1995) Young children’s knowledge about
thinking. Monogr Soc Res Child Dev 60: 1–96; discussion 97–114. Available:
http://www.ncbi.nlm.nih.gov/pubmed/7877641. Accessed 2013 October 14.
60. Brosnan SF, de Waal FBM (2003) Monkeys reject unequal pay. Nature 425:
297–299. Available: http://dx.doi.org/10.1038/nature01963. Accessed 2013
October 11.
61. Judge PG, Bruno S (2012) Transport of Functionally Appropriate Tools by
Capuchin Monkeys (Cebus apella). Am J Primatol 74: 199–209. Available:
http://doi.wiley.com/10.1002/ajp.21987. Accessed 2013 October 11.
62. de Waal FBM, Suchak M (2010) Prosocial primates: selfish and unselfish
motivations. Philos Trans R Soc Lond B Biol Sci 365: 2711–2722. Available:
http://www.ncbi.nlm.nih.gov/pubmed/20679114. Accessed 2010 September 2.
63. Pfeiffer T, Rutte C, Killingback T, Taborsky M, Bonhoeffer S (2005) Evolution of
cooperation by generalized reciprocity. Proc Biol Sci 272: 1115–1120. Available:
http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid = 1559812&tool = pmcentrez
&rendertype = abstract. Accessed 2012 March 6.
7 January 2014 | Volume 9 | Issue 1 | e87035

64. Rankin DJ, Taborsky M (2009) Assortment and the evolution of generalized
reciprocity. Evolution 63: 1913–1922. Available: http://www.ncbi.nlm.nih.gov/
pubmed/19222566. Accessed 2012 March 3.
65. Ekman P (1992) An argument for basic emotions. Cogn Emot 6: 169–200.
Available: http://www.tandfonline.com/doi/abs/10.1080/
02699939208411068. Accessed 2013 September 23.
66. Johnson-Laird PN, Oatley K (1992) Basic emotions, rationality, and folk theory.
Cogn Emot 6: 201–223. Available: http://www.tandfonline.com/doi/abs/10.
1080/02699939208411069. Accessed 2013 October 11.
67. Dawkins MS (2000) Animal Minds and Animal Emotions. Integr Comp Biol 40:
883–888. Available: http://icb.oxfordjournals.org/content/40/6/883.short.
Accessed 2013 October 11.
68. Ortony A, Norman DA, Revelle W (2005) Affect and Proto-Affect in Effective
Functioning. In: Fellous J-M, Arbib MA, editors. Who needs emotions? The
brain meets the robot. New York, NY: Oxford University Press. 173–202.
69. Weiner B (1986) Attribution, emotion, and action. In: Sorrentino RM, Higgins
ET, editors. Handbook of motivation and cognition: Foundations of social
behavior I. New York, NY. 281–312.
70. Allred KKG, Mallozzi JSJ, Matsui F, Raia CPC (1997) The Influence of Anger
and Compassion on Negotiation Performance. Organ Behav Hum Decis Process
70: 175–187. Available: http://www.sciencedirect.com/science/article/pii/
S0749597897927059. Accessed 2013 October 8.
71. Cunningham MR, Steinberg J, Grev R (1980) Wanting to and having to help:
Separate motivations for positive mood and guilt-induced helping. J Pers Soc
Psychol 38: 181–192. doi:10.1037/0022-3514.38.2.181.
72. Isen AM, Levin PF (1972) Effect of feeling good on helping: Cookies and
kindness. J Pers Soc Psychol 21: 384–388. Available: http://www.ncbi.nlm.nih.
gov/pubmed/5060754. Accessed 2013 October 11.
73. Barsade SSG (2002) The Ripple Effect: Emotional Contagion and Its Influence
on Group Behavior. Adm Sci Q 47: 644. Available: http://asq.sagepub.com/
content/47/4/644.short. Accessed 2013 September 25.
PLOS ONE | www.plosone.org
Give What You Get
74. Aureli F (2002) Conflict resolution following aggression in gregarious animals: A
predictive framework. Anim Behav 64: 325–343. Available: http://linkinghub.
elsevier.com/retrieve/doi/10.1006/anbe.2002.3071. Accessed 2011 June 16.
75. Gustison ML, MacLarnon A, Wiper S, Semple S (2012) An experimental study
of behavioural coping strategies in free-ranging female Barbary macaques
(Macaca sylvanus). Stress 15: 608–617. Available: http://informahealthcare.
com/doi/abs/10.3109/10253890.2012.668589. Accessed 2013 October 11.
76. Virgin CE, Sapolsky RM (1997) Styles of male social behavior and their
endocrine correlates among low-ranking baboons. Am J Primatol 42: 25–39.
Available: http://www.ncbi.nlm.nih.gov/pubmed/9108969. Accessed 2013
October 11.
77. Nowak MA, Sigmund K (1998) The Dynamics of Indirect Reciprocity. J Theor
Biol 194: 561–574. Available: http://www.sciencedirect.com/science/article/
pii/S0022519398907755. Accessed 2013 October 11.
78. Panchanathan K, Boyd R (2004) Indirect reciprocity can stabilize cooperation
without the second-order free rider problem. Nature 432: 499–502. Available:
http://dx.doi.org/10.1038/nature02978. Accessed 2013 September 22.
79. Piazza J, Bering JM (2008) Concerns about reputation via gossip promote
generous allocations in an economic game. Evol Hum Behav 29: 172–178.
Available: http://linkinghub.elsevier.com/retrieve/pii/S1090513807001390.
Accessed 2010 December 26.
80. Andreoni J (1990) Impure Altruism and Donations to Public Goods: A Theory of
Warm-Glow Giving. Econ J 100: 464. Available: http://www.jstor.org/stable/
2234133?origin = crossref.
81. Andreoni J, Petrie R (2004) Public goods experiments without confidentiality: A
glimpse into fund-raising. J Public Econ 88: 1605–1623. Available: http://
linkinghub.elsevier.com/retrieve/pii/S0047272703000409. Accessed 2011 Feb-
ruary 1.
82. Dana J, Weber RA, Kuang JX (2006) Exploiting moral wiggle room:
Experiments demonstrating an illusory preference for fairness. Econ Theory
33: 67–80. Available: http://link.springer.com/10.1007/s00199-006-0153-z.
Accessed 2013 October 2.
8 January 2014 | Volume 9 | Issue 1 | e87035